Echinoderms (Phylum Echinodermata) are a phylum of marine animals. The adults are recognized easily by their (usually five-point) radial symmetry, and include such well known animals as sea stars, sea urchins, sand dollars, and sea cucumbers. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone. The phylum contains about 70,000 living species,[1] making it the second-largest grouping of deuterostomes (a superphylum), after the chordates (which include the vertebrates, such as humans, sharks and frogs). Echinoderms are also the largest phylum that has no freshwater or terrestrial (land-based) representatives.
Aside from the hard-to-classify Arkarua (a Precambrian animal with Echinoderm-like traits), the first definitive members of the phylum appeared near the start of the Cambrian period. The word "Echinoderm" is derived from the Greek ἐχινοδέρματα (echinodermata), plural of ἐχινόδερμα (echinoderma), "spiny skin" from ἐχινός (echinos), "sea-urchin", originally "hedgehog,"[2] and δέρμα (derma), "skin".[3][4]
The echinoderms are important both biologically and geologically. Biologically, there are few other groupings so abundant in the biotic desert of the deep sea, as well as the shallower oceans. The more notably distinct trait, which most echinoderms have, is their remarkable powers of regeneration of tissue, organs, limbs, and of asexual reproduction, and in some cases, complete regeneration from a single limb. Geologically, the value of echinoderms is in their ossified skeletons, which are major contributors to many limestone formations, and can provide valuable clues as to the geological environment. Further, it is held by some[citation needed][who?] that the radiation of echinoderms was responsible for the Mesozoic revolution of marine life.
Two main subdivisions of echinoderms are traditionally recognised: the more familiar motile Eleutherozoa, which encompasses the Asteroidea (sea star), Ophiuroidea (brittle stars), Echinoidea (sea urchins and sand dollars), and Holothuroidea (sea cucumbers); and the sessile Pelmatozoa, which consists of the crinoids and of the extinct blastoids and Paracrinoids. Some crinoids, however, namely the feather stars, are also highly motile (able to move themselves).
In a demonstration of its motility, this Brittle Star is upside down.
The Brittle Star starts flipping itself rightside up
The Brittle Star is now rightside up.
A fifth class of Eleutherozoa consisting of just three species, the Concentricycloidea (sea daisies), were recently[5] merged into the Asteroidea. The fossil record contains a host of other classes which do not appear to fall into any extant crown group.
Echinoderms evolved from animals with bilateral symmetry. Although adult echinoderms possess pentaradial, or five-sided, symmetry, echinoderm larvae are ciliated, free-swimming organisms that organize in bilateral symmetry which makes them look like embryonic chordates. Later, the left side of the body grows at the expense of the right side, which is eventually absorbed. The left side then grows in a pentaradially symmetric fashion, in which the body is arranged in five parts around a central axis.
All echinoderms exhibit fivefold radial symmetry in portions of their body at some stage of life, even if they have secondary bilateral symmetry. Many crinoids and some seastars exhibit symmetry in multiples of the basic five, with seastars such as Helicoilaster known to possess up to 50 arms, and the sea-lily Comanthina schlegelii boasting 200.
Echinoderms have a mesodermal skeleton composed of calcareous plates or ossicles. Despite the robustness of the individual skeletal modules, complete echinoderm skeletons are rare in the fossil record. This is because they quickly disarticulate (disconnect from each other) once the encompassing skin rots away, and in the absence of tissue there is nothing to hold the plates together. The modular construction is a result of the growth system employed by echinoderms, which adds new segments at the centre of the radial limbs, pushing the existing plates outwards, as if they are being "stacked". The spines of sea urchins are most readily lost, as each spine can be moved individually and is only loosely attached in life. A walk above a rocky shore will often reveal a large number of spineless but otherwise complete sea urchin skeletons.
Skeletal elements are also deployed in some specialized ways, such as the "Aristotle's lantern" of sea urchins, crinoids' stalks, and the supportive "lime ring" of sea cucumbers.
The epidermis (skin) itself consists of cells responsible for the support and maintenance of the skeleton, as well as pigment cells, mechanoreceptor cells (which detect motion on the animal's surface), and sometimes gland cells which secrete sticky fluids or even toxins.
Echinoderms exhibit a wide range of colours.
The varied and often vivid colors of echinoderms are produced by the action of skin pigment cells. These may be light sensitive, and as a result many species change appearance completely as night falls. The reaction can happen very quickly — the sea urchin Centrostephanus longispinus changes from jet black to grey-brown in just 50 minutes when exposed to light. The colours are produced by a variable combination of colored pigments, such as the dark melanin, red carotinoids, and carotin proteins, which can be blue, green, or violet.
Echinoderms possess a unique water vascular or ambulacral (appendages such as "tube feet") system. This is a network of fluid-filled canals that function in gas exchange, feeding, and secondarily in locomotion. This system is derived from both the hydrocele and axocele. This system may have allowed echinoderms to feed without the pharyngeal slits found in other deuterostomes.
The system comprises a central ring, the hydrocele, and radial ambulacra (not to be confused with ambulacral) stretching along the body or arms. As well as assisting with the distribution of nutrients throughout the animal, the system is most obviously expressed in the tube-feet of most echinoderms. These are extensions of the water vascular system which poke through holes in the skeleton and can be extended or contracted by the redistribution of fluid between the foot and internal sac.
In the crinoids, the tube feet waft food particles captured on the radial limbs towards the central mouth; in the asteroids, the same wafting motion is employed to move the animal across the ground. Sea urchins use their feet to prevent the larvae of encrusting organisms from settling on their surfaces; potential settlers are moved to the urchin's mouth and eaten. Some burrowing sea stars poke their tube feet through the surface of the sand or mud above them into the water column and use them to attain oxygen from the water column.
Although echinoderms possess a complete digestive gut, it is very simple, often simply leading directly from mouth to anus. It can generally be divided into a pharynx, stomach, intestine and anus or cloaca.
Echinoderms also have a haemal system, and often also a perihaemal system. Both are derived from the coelom, and form an open and reduced circulatory system. This usually consists of a central ring and five radial vessels, although there is no true heart, and the blood often lacks any respiratory pigment.
Gaseous exchange occurs by dermal branchae or papulae in seastars, peristominal gills in sea urchins, genitial bursae in brittle stars and cloacal trees in holothurians. Exchange of gases also takes place through tube feet.
Echinoderms lack specialized excretory (waste disposal) organs and so nitrogenous waste, chiefly in the form of ammonia diffuses out through the respiratory surfaces.
They have a simple radial nervous system that consists of a modified nerve net — interconnected neurons with no central brain (although some do possess ganglia). Nerves radiate from central rings around the mouth into each arm or along the body; the branches of these nerves coordinate the movements of the organism.
The gonads occupy the entire body cavities of sea urchins and sea cucumbers, while the less voluminous crinoids, brittle stars, and seastars have two gonads per arm. While the primitive condition is considered to be one genital aperture, many organisms have multiple holes through which eggs or sperm may be released.
Many echinoderms have remarkable powers of regeneration. Many species routinely autotomize and regenerate arms and viscera. Sea cucumbers often discharge parts of their internal organs if they perceive danger. The discharged organs and tissues are quickly regenerated. Sea urchins are constantly replacing spines lost through damage. Sea stars (asterozoa) and Sea lillys (crinozoa) readily lose and regenerate their arms. In most cases single severed arms cannot regenerate a complete individual in the absence of at least part of the disc.[6][7][8][9] However, in a few species a single arm can survive and develop into a complete individual.[7][8][9] Some of these actively break off their arms for the purpose of asexual reproduction. During periods when they have lost the digestive tract, they live off stored nutrients and absorb dissolved organic matter directly from the water.[10]
The regeneration of lost parts involves both epimorphosis and morphallaxis. In epimorphosis stem cells - either from a reserve pool or those produced by dedifferentiation- form a blastema and generate new tissues. Morphallactic regeneration involves the movement and remodelling of existing tissues to replace lost parts. Direct transdifferentiation of one type of tissue to another during tissue replacement is also observed.[11]
The robust larval regeneration is responsible for many of them being popular model organisms in developmental biology.
Echinoderms become sexually mature after approximately two to three years, depending on the species and the environmental conditions. The eggs and sperm cells are released into open water, where fertilization takes place. The release of sperm and eggs is coordinated temporally in some species, and spatially in others. Internal fertilization has currently been observed in three species of sea star, three brittle stars and a deep water sea cucumber.
In some species of feather star, the embryos develop in special breeding bags, where the eggs are held until sperm released by a male happen to find them and fertilize the contents. This can also be found among sea urchins and sea cucumbers, where exhibit care for their young can occur, for instance in a few species of sand dollars who carry their young between the pricks of their oral side, and heart urchins possess breeding chambers. With brittle stars, special chambers can be developed near the stomach bags, in which the development of the young takes place. Species of sea cucumbers with specialized care for their offspring may also nurse the young in body cavities or on their surfaces. In rare cases, direct development without passing through a bilateral larval stage can occur in some sea stars and brittle stars.[citation needed] Another strategy that has evolved in some sea stars and brittle stars is the ability to reproduce asexually by dividing in two halves while they are small juveniles, while turning to sexual reproduction when they have reached sexual maturity.[verification needed][citation needed]
One species of seastar reproduces asexually by parthenogenesis.[12] In certain other asterozoans the adult organisms reproduce asexually for a while before they mature and reproduce sexually. In most of these species, they reproduce by transverse fission- with the disc splitting in two. Regrowth of both the disc and the arms occur[9][13] giving an animal with some large arms and some small arms during the period of growth. Though in most species at least part of the disc is needed for complete regeneration, in a few species of sea stars a single severed arm can grow into a complete individual over a period of several months.[7][8][9] In at least some of these species, they actively use this as a method of asexual reproduction.[7][14] A fracture develops on the lower surface of the arm and the arm pulls itself free from the body which holds onto the substrate during the process.[14] During the period of regrowth, they have a few tiny arms and one large arm earning them the name of "comet forms".[8][14]
Asexual reproduction by fission has also been observed in adult holothuroidea (Sea cucumbers).[15] They divide into two at a point closer to the anterior end from the middle. The two parts regenerate the missing organs over period of a few months.(see architomy)
The larvae of some echinoderm species are capable of asexual reproduction. These species belong to four of the major classes of echinoderms except crinozoans (as of 2011).[16] Asexual reproduction in the planktonic larvae occurs through numerous modes. They may autotomise parts that develop into secondary larvae, grow buds or undergo paratomy. The parts that are autotomised or the buds may develop directly into fully formed larvae or may develop through a gastrula or even a blastula stage. The parts that develop into the new larvae vary from the preoral hood (a mound like structure above the mouth), the side body wall, the postero-lateral arms or their rear ends.[16][17][18]
The process of cloning costs the larva both in resources as well as in development time. They have been observed to undergo this process when food is plentiful[19] or temperature conditions are right.[18] It has also been proposed that cloning may occur to make use of the tissues that are normally lost in metamorphosis.[20] Recent research has shown that the larvae of some sand dollars clone themselves when they detect predators (by sensing dissolved fish mucus).[18][20] Asexual reproduction produces many smaller larvae that escape planktivorous fish better.[21]
The "pluteus larva" of a sea urchin
The development of an echinoderm begins with a bilaterally symmetrical embryo, with a coeloblastula developing first. Gastrulation marks the opening of the "second mouth" that places them within the deuterostomes, and the mesoderm, which will host the skeleton, migrates inwards. The secondary body cavity, the coelom, forms by the partitioning of three body cavities.
Upon metamorphosis, each taxon produces a distinct planktonic larva, which varies in shape among the classes.[22][23] Larval stages with prominent "arms" are often referred to as pluteus larvae (often with a prefix to denote taxon).[24]
The left hand side of the larva develops into the adult organism while the right hand side eventually being absorbed; the left hand side typically becomes the oral plate.
Echinoderms are globally distributed in almost all depths, latitudes and environments in the ocean. They reach highest diversity in reef environments but are also widespread on shallow shores, around the poles — refugia where crinoids are at their most abundant — and throughout the deep ocean, where bottom-dwelling and burrowing sea cucumbers are common — sometimes accounting for up to 90 % of organisms. While almost all echinoderms are benthic — that is, they live on the sea floor — some sea-lilies can swim at great velocity for brief periods of time, and a few deep-sea sea cucumbers are fully floating. Some crinoids are pseudo-planktonic, attaching themselves to floating logs and debris, although this behaviour was exercised most extensively in the Paleozoic, before competition from such organisms as barnacles restricted the extent of the behaviour. Some sea cucumbers employ a similar strategy, hitching lifts by attaching to the sides of fish.
The larvæ of echinoderms, especially starfish and sea urchins, are pelagic, and with the aid of ocean currents can swim great distances, reinforcing the global distribution of the phylum.
The modes of feeding vary greatly between the constituent taxa. Crinoids and some brittle stars tend to be passive filter-feeders, absorbing suspended particles from passing water; sea urchins are grazers, sea cucumbers deposit feeders, and seastars are active hunters.
Crinoids employ a large net-like structure to sieve water as it is swept by currents, and to absorb any particles of matter sinking from the ocean overhead. Once a particle touches the arms of the creature, the tube feet act to swish it to the central mouth of the crinoid, where it is ingested, nutrients removed, and the remains egested through its anus to the underlying water column.
Many sea urchins graze on the surfaces of rocks, scraping off the thin layer of algae covering the surfaces. Other toothless breeds devour smaller organisms, which they may catch with their tube feet, whole. Sand dollars may perform suspension feeding.
Sea cucumbers may be suspension feeders, sucking vast quantities of sea water through their guts and absorbing any useful matter. Others use their feeding apparatus to actively capture food from the sea floor. Yet others deploy their feeding apparatus as a net, in which smaller organisms become ensnared.
While some sea stars are detritovores, extracting the organic material from mud, and others mimic the crinoids' filter feeding, most are active hunters, attacking other sea stars or shellfish. The latter are seized and held by the tube feet; sea stars then stiffen their legs, expanding the shell. The sea stars can use connective tissue to lock their arms in place and maintain a force on the prey while exerting minimal effort; the unfortunate victim must expend energy resisting the force with its adductor muscle. When the adductor tires, the sea star can insert its stomach through the opening and release gastric juices, digesting the prey alive.
Despite their low nutrition value and the abundance of indigestible calcite, echinoderms are the prey of many organisms, such as crabs, sharks, sea birds and larger starfish. Defensive strategies employed include the presence of spines, toxins, which can be inherent or delivered through the tube feet, and the discharge of sticky entangling threads by sea cucumbers. Being stabbed by a sea urchin may result in painful injury.
Echinoderms provide a key ecological role in ecosystems. For example, the grazing of sea urchins reduces the rate of colonization of bare rock; the burrowing of sand dollars and sea cucumbers depleted the sea floor of nutrients and encouraged deeper penetration of the sea floor, increasing the depth to which oxygenation occurs and allowing a more complex ecological tiering to develop. Starfish and brittle stars prevent the growth of algal mats on coral reefs, which would obstruct the filter-feeding constituent organisms. Some sea urchins can bore into solid rock; this bioerosion can destabilise rock faces and release nutrients into the ocean.
The echinoderms are also the staple diet of many organisms, most notably the otter; conversely, many sea cucumbers provide a habitat for parasites, including crabs, worms and snails. The extinction of large quantities of echinoderms appears to have caused a subsequent overrunning of ecosystems by seaweed, or the destruction of an entire reef.
Early Echinoderms (?)
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Axis scale: millions of years ago.
The first universally accepted echinoderms appear in the Lower Cambrian period (Paul and Smith 1984). Echinoderms left behind an extensive fossil record. Despite this, there are numerous conflicting hypotheses on their phylogeny. Based on their bilateral larvae, many zoologists argue that echinoderm ancestors were bilateral and that their coelom had three pairs of spaces (trimeric).
Some have proposed that radial symmetry arose in a free-moving echinoderm ancestor and that sessile groups were derived several times independently from free-moving ancestors. Unfortunately, this view does not address the significance of radial symmetry as an adaptation for a sessile existence.
The more traditional view is that the first echinoderms were sessile, became radial as an adaptation to that existence, and then gave rise to free-moving groups. This view perceives the evolution of endoskeletal plates with stereom[25] structure and of external ciliary grooves for feeding as early echinoderm developments.
The extinct members of paraphyletic Homalozoa, commonly referred to as carpoids, had stereom ossicles but were not radially symmetrical, and the status of their water-vascular system is not known. Further, extinct members of the Class Helicoplacoidea possessed three, true ambulacral grooves, and their mouth was on the side of their body.
Attachment to a substratum would have selected for radial symmetry and may have marked the origin of the Class Crinoidea. Members of Crinoidea, along with the extinct members of Class Cystoidea, were primitively attached to a substratum by an aboral stalk. An ancestor that became free-moving might have given rise to Asteroidea, Ophiuroidia, Holothuroidea, and Echinoidea.
Echinoderms sometimes pose a health threat to humans. The fine structure of the spines of certain species of sea urchins means that if the spine pierces the flesh, it may break off when an attempt is made to remove it. It may require patience — or the assistance of a physician — to fully remove the remaining piece of spine.
Echinoderms are also elements of many cuisines. Around 50,000 tons of sea urchins are captured each year, the gonads of which are consumed particularly in Japan, Peru, Spain and France. The taste is described as soft and melting, like a mix of seafood and fruit. The quality depends on the color, which can range from light yellow to bright orange.
Sea cucumbers are also considered a delicacy in some countries of south east Asia; particularly popular are the (Pineapple) roller Thelenota ananas (susuhan) and the red Halodeima edulis. They are well known as bêche de mer or Trepang in China and Indonesia. The sea cucumbers are dried, and the potentially poisonous entrails removed. The strong poisons of the sea cucumbers are often psychoactive, but their effects are not well studied. It does appear that some sea cucumber toxins restrain the growth rate of tumour cells, which has sparked interest from cancer researchers.
The calcareous tests or shells of echinoderms are used as a source of lime by farmers in areas where limestone is unavailable; indeed, 4,000 tons of the animals are used annually for this purpose. This trade is often carried out in conjunction with shellfish farmers, for whom the starfish pose a major irritation by eating their stocks.
Sea-urchin and sand dollar skeletons are popular collectibles, as are dried starfish.
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- ^ a b c Monks, Sarah P. (1904-04-01). "Variability and Autotomy of Phataria". Proceedings of the Academy of Natural Sciences of Philadelphia 56 (2): 596–600. ISSN 0097-3157. JSTOR 4063000.
- ^ Kille, Frank R. (1942). "Regeneration of the Reproductive System Following Binary Fission in the Sea-Cucumber, Holothuria parvula (Selenka)". Biological Bulletin 83 (1): 55–66. DOI:10.2307/1538013. ISSN 0006-3185. JSTOR 1538013. http://www.biolbull.org/cgi/reprint/83/1/55.pdf. Retrieved 2011-07-15.
- ^ a b Eaves, Alexandra A.; A. Richard Palmer (2003). "Reproduction: Widespread cloning in echinoderm larvae". Nature 425 (6954): 146. DOI:10.1038/425146a. ISSN 0028-0836.
- ^ Jaeckle, William B. (1994-02-01). "Multiple Modes of Asexual Reproduction by Tropical and Subtropical Sea Star Larvae: An Unusual Adaptation for Genet Dispersal and Survival". Biological Bulletin 186 (1): 62–71. DOI:10.2307/1542036. ISSN 0006-3185. JSTOR 1542036. http://www.biolbull.org/cgi/reprint/186/1/62.pdf. Retrieved 2011-07-13.
- ^ a b c Vaughn, Dawn (2009-10-01). "Predator-Induced Larval Cloning in the Sand Dollar Dendraster excentricus: Might Mothers Matter?". Biol Bull 217 (2): 103–114. http://www.biolbull.org/cgi/content/abstract/217/2/103. Retrieved 2011-07-16.
- ^ McDonald, Kathryn A.; Dawn Vaughn (2010-08-01). "Abrupt Change in Food Environment Induces Cloning in Plutei of Dendraster excentricus". Biol Bull 219 (1): 38–49. http://www.biolbull.org/cgi/content/abstract/219/1/38. Retrieved 2011-07-16.
- ^ a b Vaughn, Dawn; Richard R. Strathmann (2008-03-14). "Predators Induce Cloning in Echinoderm Larvae" (Free registration required for full text.). Science 319 (5869): 1503. DOI:10.1126/science.1151995. JSTOR 40284699. PMID 18339931. http://www.sciencemag.org/content/319/5869/1503.abstract. Retrieved 2011-07-16.
- ^ Vaughn, Dawn (2010-03). "Why run and hide when you can divide? Evidence for larval cloning and reduced larval size as an adaptive inducible defense". Marine Biology 157 (6): 1301–1312. DOI:10.1007/s00227-010-1410-z. ISSN 0025-3162. http://www.springerlink.com/content/026u662661533477/. Retrieved 2011-07-16.
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- This article incorporates information from this version of the equivalent article on the German Wikipedia.
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